Journal of the Royal Society of Western Australia, 81:177-182, 1998

New species of and (freshwater Copepoda: ) from Western Australia and Queensland

I A E Bayly

Department of Biological Sciences, Monash University Clayton, Vic 3168

Manuscript received September 1997; accepted March 1998

Abstract A small calanoid , Calamoecia halsei sp nov, is described from shallow, turbid waters in claypans of the Carnarvon region of Western Australia. The genus Calamoecia Brady, which is endemic to the Australasian region, now contains 15 species. A larger calanoid copepod, Boeckella timmsi sp nov, is described from a shallow turbid claypan in Currawinya National Park located in Queensland just north of the border with New South Wales. The genus Boeckella de Guerne and Richard (which also occurs on sub-Antarctic islands, Antarctica and South America) now contains 21 Australasian species.

Introduction contrast illumination) and drawn using a Wild M20 microscope fitted with a Treffenberg drawing tube. The Most calanoid occurring in Australian system of abbreviations given in full by Bayly (1992a,b) inland waters are in the genera Calamoecia and Boeckella. was used in the description of the first and fifth legs. As a generalization, the smaller calanoids (body length less than ca 1.4 mm, but C. attenuata is longer) belong to Calamoecia while the larger ones (female body length range ca 1.4 – 4.5 mm, but some including B. minuta are shorter) belong to Boeckella. Eight species belonging to Family Giesbrecht those genera were described in the late nineteenth century and early twentieth century by the Norwegian Genus Calamoecia Brady carcinologist G O Sars, in some cases after rearing them in Oslo from resting eggs contained in parcels of dried Calamoecia halsei sp nov mud consigned from Australia. The latter were part of an (Fig 1A–G) historic series of hatching experiments which resulted in the description of a significant portion of the freshwater Specimens examined lower fauna (Branchiopoda, Cladocera, WESTERN AUSTRALIA (Carnarvon region): Chagra Ostracoda and Copepoda) of Australia. Boeckella minuta Well claypan, Jimba Jimba Station, 25° 11.78' S, 114° 57.04' was described in this manner in 1896 (Sars 1896). E, 22.viii.1994, 10 female (mean length prosome 0.94 mm, Calamoecia was revised by Bayly (1961, 1962a) who mean length to end of caudal rami 1.35 mm), 10 male recognised 12 species, to which were added C. elongata (mean lengths 0.73 mm, 1.04 mm); un-named claypan, (Bayly 1979) and C. zeidleri (Bayly 1984). Boeckella was Jimba Jimba Station, 25° 04.29' S, 115° 03.50' E, revised by Bayly (1964) with the recognition of 17 22.viii.1994, 10 female (mean lengths 0.97 mm, 1.37 mm), Australasian species, to which were added B. bispinosa 10 male (mean lengths 0.76 mm, 1.08 mm); un-named (Bayly 1967), a New Zealand endemic B. tanea (Chapman canegrass pan, Wooramel Station, 25° 40.87' S, 114° 13.23' E, 1973), and B. shieli (Bayly 1985). A comprehensive key to 24.viii.1994, 2 female (mean length prosome 0.92 mm), 4 these two genera was published by Bayly (1992a). Now, male (mean lengths 0.73 mm, 0.99 mm); un-named after more than a decade without addition to either claypan, Coolcalalaya Station, 27° 31.49' S, 115° 05.24' E, genus, a new species of both has come to hand at 30.viii.1994, 7 female (mean length prosome 0.89 mm), 2 practically the same time in 1997 and the two are male (mean lengths 0.71 mm, 0.95 mm); ephermeral described below. marsh, Brickhouse Station, 24° 57.85' S, 113° 42.27' E, 25.viii.1994, 1 male. All specimens collected by S A Halse.

Methods Type material: Holotype male, allotype female, paratypes 30 male, 30 female. Holotype and allotype Specimens were measured under a Wild M7 stereo- mounted on microslides, paratypes unmounted in vial. microscope fitted with an eyepiece micrometer and Western Australian Museum Crustacean Department dissected with tungsten needles in PVA-lactophenol registration numbers WAM 639–97 to 641–97. Type mountant on a microslide under the same microscope. locality: Chagra Well claypan, 25° 11.78' S, 114° 57.04' E. Appendages were examined (with bright field and phase- Description of male Size: Length of prosome 0.68 – 0.81 mm, length to end of © Royal Society of Western Australia 1998 caudal rami 0.90 – 1.13 mm.

177 Journal of the Royal Society of Western Australia, 81(4), December 1998

A B C

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D E F G

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Figure 1. Calamoecia halsei sp nov; A, leg of first pair (drawn from male but identical in female except for larger size) [see text regarding arrows]; B & C, posterior and anterior aspects, respectively, of male fifth legs [arrows point to a unique, diagnostic feature]; D, leg of female fifth pair, showing whole of exopodite; E, as for D but showing whole of endopodite [arrows in D and E point to unusually long process from middle exopodite segments]; F, ventral aspect of female genital segment [arrow points to distinctive ridge and furrow on right side]; G, left lateral aspect of female genital segment (position of ridge and furrow located mid-laterally on right side shown by dotted crescent).

178 I A E Bayly: New species of Calamoecia and Boeckella copepods

First legs (Fig 1A): Outer edge spines on Re1 and Re3 a) the right Re claw is without an inner spine (see arrows Fig 1A) unusual within freshwater b) the right Ri is 2-segmented with Ri2 exceeding Ri1 Centropagidae in having abnormally large secondary in length by at least three times spinules. [These elaborate outer spines not occurring on exopods of legs 2–4]. c) the left Ri is 1-segmented, very elongated, considerably exceeds the left Re in length, and Fifth legs (Fig 1B,C): Right Re claw bent through almost bears five spines of which four are terminal and a right angle, lacking an inner spine. Right Ri 2- one sub-terminal on the outside. segmented with Ri2 four times as long as Ri1; Ri2 with upraised subrectangular thickening near proximal outer Although I have drawn attention to the unusually corner on anterior face (unique feature – see arrow Fig elaborate spines on the outer edge of the exopods of the 1B,C) and typically with two spines at extremity – a large first pair of legs (Fig 1A) of C. halsei, it should be noted outer curved hook or claw and a smaller inner straight or that these appendages have not been described for all slightly curved spine [extremity sometimes (observed species of Calamoecia. However, it can be said one specimen in 15 examined) with third small, straight confidently that they do not occur in all species; they inner spine]. Left Re 2-segmented; Re2 spatulate or are absent, for example, in C. lucasi, the type species of spoon-shaped (often bent out of alignment shown in Fig the genus (New Zealand material of this species 1B,C because of fragile nature) with smoothly rounded examined for this feature by the author in July 1997), C. extremity marked with minute pits or dots, and with seta australica (Sars 1908, Plate ii, Fig 13) and C. tasmanica inserted little short of half-way along outer edge. Left Ri [see drawing by Sars (1912, Plate vii, Fig 9) of Brunella 1-segmented, very elongated and exceeding left Re in longicornis, which is a junior synonym of C. tasmanica]. length, typically armed with five spines of which four The latter drawing of Sars also shows no outer spine on are terminal (second from inside shortest with remainder Re1 (formula for outer exopod spines 0.0.2) whereas this sub-equal) and one sub-terminal on outer edge [sixth spine is present in most species of Calamoecia spine sometimes (observed one specimen in 15 (corresponding formula 1.0.2; Fig 1A). examined) present on inner edge, short distance from Most of the material examined here came from very extremity]. shallow, highly turbid (turbidity 10,000–36,000 NTU) pans that contained decidedly fresh water (TDS 190–550 Description of female -1 -1 mg L , K25 140–920 µS cm ). The ephemeral marsh was a Size: Length of prosome 0.81 – 1.00 mm, length to end of little less turbid (9,100 NTU) and slightly higher in electrolytes (K 1,110 µS cm-1) but still fresh. (Chemical caudal rami 1.13 – 1.45 mm. 25 data from S A Halse, CALM, unpublished observations.) First legs: Outer edge spines on Re1 and Re3 as described for male. Derivation of specific name: Named for Western Australian limnologist S A Halse, who collected all the Fifth legs (Fig 1D,E): Outgrowth from inner distal material of this new species and drew it to my corner of Re2 (the structural feature that is of key attention. importance in defining the family Centropagidae) long, slender and only slightly curved, extending almost to extremity of long terminal spine on Re3 (and sometimes Family Centropagidae Giesbrecht beyond). In several species of Calamoecia this outgrowth does not reach past the end of Re3 excluding its Genus Calamoecia Brady terminal spine; cf C. gibbosa where this structure is relatively massive and strongly curved (Bayly 1979, Fig Calamoecia cf lucasi Brady 3E,F). Re3 typically with 5 spines (3 subequal on inner edge, 1 long terminal and 1 short on outer edge) but (Gascoyne-Murchison form) sometimes (1 in 12 legs examined) with only 4 spines (2 only subequal ones on inner edge). Ri 1-segmented, Calamoecia lucasi Brady. The Cue population, Bayly, with a total of 7 setae (4 on inner edge, 1 terminal and 2 1984, pp 149–150, Figs 4A–D. on outer edge) plus relatively short, thin spine Specimens examined immediately to inside of terminal seta. WESTERN AUSTRALIA (Carnavon region): Dywers Urosome: Genital segment (Fig 1F,G) about 1.3 times as Pan, 24 ° 26’ 00” S, 114° 27’ 18” E, 19.viii.1994, 10 females long as maximum width viewed ventrally or dorsally, and 10 males, collected S A Halse. with thickened ridge and indentation mid-laterally on right side about one third length of segment from Preliminary reassessment of taxonomic status posterior edge (see arrow, Fig 1F). Bayly (1984) referred a form of Calamoecia collected Remarks: If one attempts to key out this species using from a pond near Cue to C. lucasi, In doing so, I pointed the dichotomous key given by Bayly (1992a) the out that the Cue form differed significantly from following key numbers would apply; 1, 2, 3, 5 and 7. populations in eastern Australia and New Zealand in Dichotomy number seven accommodates C. australica several features involving secondary sexual structures. I Sars 1908 and C. canberra Bayly 1962, both of which are now find that the above Carnarvon material, which distinctly different from C. halsei. In terms of secondary agrees exactly with the Cue population in the male and sexual structures C. halsei is most closely related to C. female fifth legs and the female genital segment, can be zeidleri, C. australica and C. canberra. This group of four clearly distinguished from C. lucasi sensu stricto, by species share the following features in the male fifth legs; structural features that are not of a secondary sexual

179 Journal of the Royal Society of Western Australia, 81(4), December 1998 nature. On the terminal exopodite segments of the first species of Boeckella, but does occur as an invariable pair of legs of the Western Australian forms, the two feature in some South American Boeckella such as B. outer edge spines each have three or four large calcaris and B. palustris). secondary spinules along both edges, and the outer edge of the stoutest terminal spine has more than 25 large Description of female secondary spinules. In brief, these three spines are closely similar to, if not identical with, those in C. halsei sp nov Size: Length of prosome measured mid-dorsally 1.42 – (cf Fig 1A). In C. lucasi sensu stricto, however, these three 1.49 mm, length to end of caudal rami 1.91 – 2.03 mm. spines have more numerous and extremely fine hair-like General body proportions (Fig 2D): More robust than outgrowths along their edges. I now think it very most limnetic congeners; prosome 2.2 times as long mid- probable that the “Gascoyne-Murchison form of C. lucasi” dorsally as maximum width (viewed dorsally or is, in fact, a new species, but I do not wish to proceed ventrally), and 2.35 times as long as urosome excluding with a formal description at this stage. caudal setae. Antennules relatively short, not reaching beyond end of prosomal “wings” when extended Family Centropagidae Giesbrecht posteriorly. Last prosomal segment produced posteriorly on both sides into large “wings” or lobes reaching to Genus Boeckella De Guerne & Richard posterior margin of urosome segment 2. Boeckella timmsi sp nov Fifth legs (Fig 2C): Re3 with 7 spines – 4 on inner edge, 1 terminal (exceeding Re3 in length) and 2 on outer edge. (Fig 2A–E) Ri 3-segmented, reaching to end of Re2 excluding inner process, setal formula 1.1.222. Specimens examined Urosome: Genital segment (Fig 2E) with highly QUEENSLAND: Claypan, near Coomburra distinctive shape, squarish, length not greater than Waterhole, Currawinya National Park, 28° 47' S, 144° 22' maximum width viewed ventrally or dorsally, with two E 24.vii.1996; 7 females (mean length prosome [measured rounded outgrowths and concavity between them on mid-dorsally to exclude large “wings” on last prosomal both sides. segment] 1.45 mm, mean length to end of caudal rami 1.98 mm), 10 males (mean length prosome [measured Remarks: If one attempts to key out this species using mid-laterally] 1.14 mm, mean length to end of caudal the dichotomous key provided by Bayly (1992a) the rami 1.51 mm). Collected by B V Timms. following key numbers would apply; 1, 2, 5, 6, 7, 8 and 9. This is a pathway that leads to B. triarticulata (Thomson), Type material : Holotype male, allotype female, B. fluvialis Henry, B. hamata Brehm (a New Zealand paratypes 13 male, 12 female. Holotype and allotype endemic) and B. robusta Sars all of which differ very mounted on microslides, paratypes unmounted in vial. significantly from B. timmsi. The latter lacks the large Queensland Museum registered numbers QMW22252–4. teeth attached to the inner distal corner of the left B2 and Type locality: Claypan 1.5 km north-east of Coomburra the relatively much longer left Re1 found in B. Waterhole, Currawinya National Park, 28° 47' S, 144° 22' E. triarticulata and B. fluvialis. The left Re1 of B. timmsi is relatively much shorter than that in B. hamata and B. Description of male robusta, being more comparable with that in B. symmetrica Size: Length of prosome measured mid-laterally 1.00– Sars. 1.16 mm, length to end of caudal rami 1.45–1.62 mm. The question of the taxonomic value of the size and shape of any posterior “wings” developed from the last Fifth legs (Fig 2A,B): Right Ri sub-triangular, attached prosomal segment of female boeckellas is a somewhat more proximally on inner edge of right B2 than is vexed one. In some limnetic species they seem not to typically the case, 2-segmented (Fig 2A) or 3-segmented occur and in others, such as B. propinqua Sars, they are (Fig 2B) with the second, or more distal, line of clearly subject to a large amount of intraspecific variation segmentation being often weak or absent, proximal inner (cf Bayly 1962b, Fig 18). On the other hand, in species projection commonly somewhat differentiated from associated with shallow bodies of temporary water such remainder of Ri (Fig 2A), distal extremity sharply as B. saycei Sars, B. nyoraensis Searle and B. bispinosa Bayly pointed (Fig 2B) or bluntly rounded (Fig 2A) reaching these lobes seem always to be large and of some just less than half-way along inner edge of right Re2. Left taxonomic importance (cf Bayly 1979, Figs 5C and 6A,C). B2 with projection at inner distal corner reaching to end The large prosomal lobes of B. timmsi and the shallow, of left Ri1 and armed with 10–20 very fine teeth. Left Ri temporary nature of its habitat are in agreement with 2-segmented with terminal segment (Ri2) sub-circular this pattern. The extreme shallowness of its habitat, and extending about 0.6 times distance along inner edge together with its robust body form and relatively short of left Re1. Left Re1 short relative to right Re, distal antennules, suggest that this species may be extremity lying directly opposite a point only 0.4–0.5 fundamentally littoral or benthic in its habits (cf Timms times distance along inner edge of right Re2. Left Re2 1979). This species came from a large (4 ha), very shallow (proximal portion of left Re claw) with prominent outer (< 40 cm deep) and highly turbid (turbidity 6000 FTU) spine inserted at 0.25 times distance along segment and claypan that contained fresh water (K 430 µS cm-1). commonly (about one-third of population; Fig. 2B) with 25 second (much smaller) outer spine inserted at 0.75 times Derivation of specific name: Named for noted distance along segment (this second outer spine on the Australian limnologist B V Timms, who collected this left Re2 is not a normal feature of any other Australian new species and drew it to my attention.

180 I A E Bayly: New species of Calamoecia and Boeckella copepods 0.1 mm

0.2 mm 0.1 mm 0.5 mm

Figure 2. Boeckella timmsi sp nov; A & B, posterior and anterior aspects, respectively, of male fifth legs; C, leg of female fifth pair; D, dorsal aspect of female (excluding most of antennules); E, ventral aspect of female genital segment (plus urosome segment 2).

181 Journal of the Royal Society of Western Australia, 81(4), December 1998

Acknowledgments: I thank S A Halse and B V Timms for providing me Bayly I A E 1984 A new species of Calamoecia (Copepoda: with the material on which this paper is based. It is to the credit of both Calanoida).from South Australia, and comments on three of them that, on the basis of their own preliminary observations, they congeners. Transactions of the Royal Society of South thought that they were in possession of a new species. I am indebted to C Australia 108:147–154. Petersen and C Logan for producing the typescript. Bayly I A E 1985 A new species of Boeckella (Copepoda: Calanoida). from Western Australia, and comments on two other congeners. Records of the Western Australian Museum References 12:79–84. Bayly I A E 1992a The non-marine Centropagidae (Copepoda: Bayly, I A E 1961 A revision of the inland water genus Calanoida) of the World [Guides to the identification of the Calamoecia (Copepoda: Calanoida). Australian Journal of Microinvertebrates of the Continental Waters of the World Marine and Freshwater Research 12:54–91. No. 2]. SPB Academic Publishing, The Hague. Bayly I A E 1962a Additions to the inland water genus Bayly I A E 1992b Fusion of the genera Boeckella and Calamoecia (Copepoda: Calanoida). Australian Journal of Pseudoboeckella (Copepoda) and revision of their species from Marine and Freshwater Research 13:252–264. South America and sub-Antarctic islands. Revista Chilena de Bayly, I A E 1962b Ecological studies on New Zealand lacustrine Historia Natural 65:17–63. zooplankton with special reference to Boeckella propinqua Sars Chapman M A 1973 A new species of Boeckella (Copepoda: (Copepoda: Calanoida). Australian Journal of Marine and Calanoida) from Northland, New Zealand. New Zealand Freshwater Research 13:143–197. Journal of Marine and Freshwater Research 7:153–157. Bayly I A E 1964 A revision of the Australasian species of the Sars G O 1896 On freshwater Entomostraca from the freshwater genera Boeckella and Hemiboeckella (Copepoda: neighbourhood of Sydney, partly raised from dried mud. Calanoida). Australian Journal of Marine and Freshwater Archiv for Mathematik og Naturvidenskab 18(3):1–81. Research 15:180–238. Sars G O 1908 Fresh-water Copepoda from Victoria, southern Bayly I A E 1967 A new species of Boeckella (Copepoda: Australia. Archiv for Mathematik og Naturvidenskab Calanoida) and additional comments on some other species 29(7):3–24. of the genus. Papers and Proceedings of the Royal Society of Sars G O 1912 Additional notes on fresh-water Calanoida from Tasmania 101:97–101. Victoria, southern Australia. Archiv for Mathematik og Bayly I A E 1979 Further contributions to a knowledge of the Naturvidenskab 32(13):3–20. centropagid genera Boeckella, Hemiboeckella and Calamoecia Timms B V 1979 Body proportions in limnetic and littoral (athalassic calanoid copepods). Australian Journal of Marine calanoid copepods. Australian Journal of Marine and and Freshwater Research 30: 103–127. Freshwater Research 30:417–419.

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